Talpidae

Talpidae; Temporal range:Late Eocene–Recent PreꞒ Ꞓ O S D C P T J K Pg N
	Left column: Top:Pyrenean desman; Bottom:European mole; ; Right column: Top:Small Japanese mole; Middle:Russian desman; Bottom:Eastern mole; ;
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Eulipotyphla
Superfamily:	Talpoidea
Family:	Talpidae; G. Fischer,1814
Type genus
	Talpa; Linnaeus,1758
Subfamilies
	Talpinae; Scalopinae; Uropsilinae

ThefamilyTalpidae^[1](/ˈtælpɪdiː/) includes themoles(some of whom are calledshrew molesanddesmans) who are smallinsectivorousmammals of theorder Eulipotyphla.Talpidsare alldigging animalsto various degrees: moles are completely subterranean animals; shrew moles andshrew-like molessomewhat less so; and desmans, while basically aquatic, excavate dry sleeping chambers; whilst the quite uniquestar-nosed moleis equally adept in the water and underground. Talpids are found across theNorthern HemisphereofEurasiaandNorth America(although none are found inIrelandnor in theAmericassouth of northernMexico), and range as far south as themontaneregions of tropicalSoutheast Asia.

The first talpids evolved fromshrew-like animals which adapted to digging late in theEocenein Europe.Eotalpa anglicais the oldest known mole, it was discovered in the Late Eocene deposits ofHampshire Basin,UK.^[2]The most primitive living talpids are believed to be theshrew-like moles,with other species having adapted further into the subterranean, and, in some cases, aquatic lifestyles.^[3]

Characteristics

Talpids are small, dark-furred animals with cylindrical bodies and hairless, tubular snouts. They range in size from the tinyshrew molesof North America, as small as 10 cm in length and weighing under 12 grams, to theRussian desman,with a body length of 18–22 cm, and a weight of about 550 grams. Thefurvaries between species, but is always dense and short; desmans have waterproof undercoats and oilyguard hairs,while the subterranean moles have short, velvety fur lacking any guard hairs. The forelimbs of moles are highly adapted for digging, with powerful claws, and the paws turned permanently outwards to aid in shovelling dirt away from the front of the body. By contrast, desmans have webbed paws with a fringe of stiff fur to aid in swimming. Moles generally have short tails, but those of desmans are elongated and flattened.^[4]

All species have small eyes and poor eyesight, but only a few are truly blind.^[4]The external ears are very small or absent.^[5]Talpids rely primarily on their sense oftouch,having sensoryvibrissaeon their faces, legs, and tails. Their flexible snouts are particularly sensitive. Desmans are able to close both their nostrils and ears while diving. Unusually, thepenisof talpids points backwards, and they have noscrotum.^[4]

Females have six or eightteats.Both sexes have claws on all five fingers and on all five toes. The paw has an additional bone called theos falciforme.In burrowing moles, theclavicleand thehumeral headare connected. Thetibiaand thefibulaare partially fused in all talpids. Thepubisdoes not connect the two halves of thepelvic girdle.Theskullis long, narrow, and rather flattened.^[5]

Talpids are generally insectivorous. Moles eatearthworms,insect larvae, and occasionally slugs, while desmans eat aquatic invertebrates such as shrimps, insect larvae, andsnails.Talpids have relatively unspecialized teeth, with thedental formula:

Dentition
2-3.1.3-4.3
1-3.0-1.3-4.3

Sex characteristics

Many species of talpid moles exhibit peniform clitorises that are tunneled by the urethra and are found to have erectile tissue, most notably species from theTalpagenus found in Europe.^[6]Unique to this clade are the presence ofovotestes,wherein the female ovary also is mostly made up of sterile testicular tissue that secretes testosterone with only a small portion of the gonad containing ovarian tissue. Genetic studies have revealed that females of these species have an XX genotype and do not have any translocated Y-linked genes.^[6]Detailed developmental studies ofTalpa occidentalishave revealed that the female gonads develop in a "testis-like pattern".DMRT1,a gene that regulates development ofSertoli cells,was found to be expressed in female germ cells before meiosis, however no Sertoli cells were present in the fully-developed ovotestes. Additionally, the female germ cells only enter meiosis postnatally, a phenomenon that has not been found in any othereutherian mammal.^[6]Phylogenetic analyses have suggested that, like in lemuroids, this trait must have evolved in a common ancestor of the clade, and has been "turned off and on" in different talpid lineages.^[7]

Female European moles are highly territorial and will not allow males in to their territory outside of breeding season, the probable cause of this behavior being the high levels of testosterone secreted by the female ovotestes.^[6]^[7]During the non-breeding season, their vaginal opening is covered by skin, akin to the condition seen in mouse and dwarf lemurs.^[6]

Behavior

Desmans and shrew moles are primarilynocturnal,but moles are active day and night, usually travelling above ground only under cover of darkness. Most moles dig permanentburrows,and subsist largely on prey that falls into them. The shrew moles dig burrows to access deep sleeping chambers, but forage for food on the forest floor by night. Desmans dig burrows in riverbanks for shelter and forage in the water of rivers and lakes. The star-nosed mole is able to make a living much as other moles do, but are also very capable aquatic creatures, where they are able to smell underwater by using their unique proboscis to hold out a bubble of air into the water.

Talpids appear to be generally quite antisocial animals, and although at least one species, the star-nosed mole, will share burrows, talpids are known to engage in much territorial behavior, including extraordinarily fast battles.^[4]

Classification

The family is divided into three subfamilies, 19 genera and 59 species.

Family Talpidae
- SubfamilyUropsilinae- Asian shrew-like moles (Chinese shrew moles)
  - GenusUropsilus- eight species in China, Bhutan, and Myanmar
    - Equivalent-teeth shrew mole,U. aequodonenia
    - Anderson's shrew mole,U. andersoni
    - Black-backed shrew mole,U. atronates
    - Dabie Mountains shrew mole,U. dabieshanensis
    - Gracile shrew mole,U. gracilis
    - Inquisitive shrew mole,U. investigator
    - Snow Mountain shrew mole,U. nivatus
    - Chinese shrew mole,U. soricipes
- SubfamilyScalopinae- New World moles
  - TribeCondylurini- condylurine moles
    - GenusCondylura- one species in eastern North America
      - Star-nosed mole,C. cristata
  - TribeScalopini- scalopine moles
    - GenusAlpiscaptulus- one species in China
      - Medog mole,A. medogensis
    - GenusParascalops- one species in northeastern North America
      - Hairy-tailed mole,P. breweri
    - GenusScalopus- one species in eastern North America
      - Eastern mole,S. aquaticus
    - GenusScapanulus- one species in China
      - Gansu mole,S. oweni
    - GenusScapanus- five species in western North America
      - Mexican mole,S. anthonyi^[8]
      - Northern broad-footed mole,S. latimanus
      - Southern broad-footed mole,S. occultus^[9]
      - Coast mole,S. orarius
      - Townsend's mole,S. townsendii
- SubfamilyTalpinae- Old World moles, desmans, and shrew moles
  - TribeTalpini- talpine moles
    - GenusEuroscaptor- ten species in East, South, and Southeast Asia
      - Greater Chinese mole,E. grandis
      - Kloss's mole,E. klossi
      - Kuznetsov's mole,E. kuznetsovi
      - Long-nosed mole,E. longirostris
      - Malaysian mole,E. malayanus^[10]
      - Himalayan mole,E. micrurus
      - Ngoc Linh mole,E. ngoclinhensis
      - Orlov's mole,E. orlovi
      - Small-toothed mole,E. parvidens
      - Vietnamese mole,E. subanura^[11]
    - GenusMogera- nine species from East Asia
      - Echigo mole,M. etigo
      - Small Japanese mole,M. imaizumii
      - Insular mole,M. insularis
      - Kano's mole,M. kanoana
      - La Touche's mole,M. latouchei
      - Ussuri mole,M. robusta
      - Sado mole,M. tokudae
      - Senkaku mole,M. uchidai
      - Japanese mole,M. wogura
    - GenusOreoscaptor- one species in Japan
      - Japanese mountain mole,O. mizura
    - GenusParascaptor- one species in southern Asia
      - White-tailed mole,P. leucura
    - GenusScaptochirus- China
      - Short-faced mole,S. moschatus
    - GenusTalpa- thirteen species, Europe and western Asia
      - Altai mole,T. altaica
      - Aquitanian mole,T. aquitania^[12]
      - Blind mole,T. caeca
      - Caucasian mole,T. caucasica
      - Père David's mole,T. davidiana
      - European mole,T. europaea
      - Levant mole,T. levantis
      - Martino's mole,T. martinorum
      - Spanish mole,T. occidentalis
      - Ognev's mole,T. ognevi
      - Roman mole,T. romana
      - Balkan mole,T. stankovici
      - Talysch mole,T. talyschensis
  - TribeScaptonychini
    - GenusScaptonyx- one species in China and Myanmar
      - Long-tailed mole,S. fusicauda
  - TribeDesmanini- desmans
    - GenusDesmana
      - Russian desman,D. moschata
    - GenusGalemys
      - Pyrenean desman,G. pyrenaicus
  - TribeUrotrichini- Japanese shrew moles
    - GenusDymecodon
      - True's shrew mole,D. pilirostris
    - GenusUrotrichus
      - Japanese shrew mole,U. talpoides
  - TribeNeurotrichini- New World shrew moles
    - GenusNeurotrichus- Pacific northwest US, southwest British Columbia
      - American shrew mole,N. gibbsii

Some studies suggest that this classification into three subfamilies is not entirely accurate, findingUropsilinaeto be the mostbasalmember, thenDesmanini,then acladecomprisingNeurotrichini,Scaptonychini,andUrotrichini,then theCondylurini,and thenTalpiniandScalopinibeingsister groupsto one another. The current classification into 3 subfamilies renders bothTalpinaeandScalopinae paraphyletic.^[13]

Unrelated mammals built like moles

The following mammals have burrowing habits, and have by virtue ofconvergent evolutionmany derived characters in common with true moles from the family Talpidae but are nonetheless unrelated.

Marsupial moles(2 species):Notoryctes typhlops,andN. caurinus.
Golden moles(21 species), belonging to theAfrotheria.

Relationship with humans

All species in the family Talpidae are classed as "prohibited new organisms" under New Zealand'sHazardous Substances and New Organisms Act 1996,preventing them from being imported into the country.^[14]

References

^Hutterer, R. (2005).Wilson, D.E.;Reeder, D.M. (eds.).Mammal Species of the World: A Taxonomic and Geographic Reference(3rd ed.). Johns Hopkins University Press. pp. 300–311.ISBN 978-0-8018-8221-0.OCLC 62265494.
^Jerry J. Hooker. Skeletal adaptations and phylogeny of the oldest mole Eotalpa (Talpidae, Lipotyphla, Mammalia) from the UK Eocene: the beginning of fossoriality in moles. The Paleontological Association. Volume59, Issue 2, March 2016. pp. 195-216.doi.org/10.1111/pala.12221
^Savage, RJG, & Long, MR (1986).Mammal Evolution: an illustrated guide.New York: Facts on File. p.53.ISBN 0-8160-1194-X.{{cite book}}:CS1 maint: multiple names: authors list (link)
^^a ^b ^c ^dGorman, Martyn (1984). Macdonald, D. (ed.).The Encyclopedia of Mammals.New York: Facts on File. pp.766–769.ISBN 0-87196-871-1.
^^a ^bGrzimek, Bernhard.Grzimek's Animal Life Encyclopedia, Volume 10: Mammals I.New York: Van Nostrand Reinhold Company, 1975. Print.
^^a ^b ^c ^d ^eJiménez, R.; Barrionuevo, F.J.; Burgos, M. (2013). "Natural Exceptions to Normal Gonad Development in Mammals".Sexual Development.7(1–3): 147–162.doi:10.1159/000338768.ISSN 1661-5433.PMID 22626995.S2CID 8721211.
^^a ^bCarmona, F. David; Motokawa, Masaharu; Tokita, Masayoshi; Tsuchiya, Kimiyuki; Jiménez, Rafael; Sánchez-Villagra, Marcelo R. (15 May 2008). "The evolution of female mole ovotestes evidences high plasticity of mammalian gonad development".Journal of Experimental Zoology Part B: Molecular and Developmental Evolution.310B(3): 259–266.doi:10.1002/jez.b.21209.ISSN 1552-5007.PMID 18085526.
^Yates, Terry L.; Jorge Salazar-Bravo (2004). "A Revision OfScapanus latimanus,with the Revalidation of a Species Of Mexican Mole ". In Sánchez-Cordero V.; Medellín R.A. (eds.).Contribuciones Mastozoológicas En Homenaje A Bernardo Villa(PDF).Instituto De Biología e Ins Tituto De Ecología, Unam, México. pp. 479–496.
^Castañeda, Sergio Ticul Alvarez; Cortés-Calva, Patrica (2021-05-09)."Revision of moles in the genus Scapanus".THERYA.12(2): 275.doi:10.12933/therya-21-1174.ISSN 2007-3364.S2CID 236583289.
^Redescription of the Malaysian Mole as to be a true speciesEuroscaptor malayana^{[permanent dead link]}
^Kawada, S-I.; et al. (2012). "A new species of mole of the genusEuroscaptor(Soricomorpha, Talpidae) from northern Vietnam ".Journal of Mammalogy.93(3): 839–850.doi:10.1644/11-MAMM-A-296.1.
^Nicolas, Violaine; Martínez-Vargas, Jessica; Hugot, Jean-Pierre (2017)."Talpa aquitania sp. nov. (Talpidae, Soricomorpha), a new mole species from SW France and N Spain"(PDF).Mammalia.81(6): 641–642.doi:10.1515/MAMMALIA-2017-0057.S2CID 90926022.
^Schwermann, Achim H.; He, Kai; Peters, Benjamin J.; Plogschties, Thorsten; Sansalone, Gabriele (2019). "Systematics and macroevolution of extant and fossil scalopine moles (Mammalia, Talpidae)".Palaeontology.62(4): 661–676.doi:10.1111/pala.12422.ISSN 1475-4983.S2CID 134096608.
^Hazardous Substances and New Organisms Act 2003 - Schedule 2 Prohibited new organisms,New Zealand Government,retrieved26 January2012

[msw3-1] Hutterer, R. (2005).Wilson, D.E.;Reeder, D.M. (eds.).Mammal Species of the World: A Taxonomic and Geographic Reference(3rd ed.). Johns Hopkins University Press. pp. 300–311.ISBN 978-0-8018-8221-0.OCLC 62265494.

[2] Jerry J. Hooker. Skeletal adaptations and phylogeny of the oldest mole Eotalpa (Talpidae, Lipotyphla, Mammalia) from the UK Eocene: the beginning of fossoriality in moles. The Paleontological Association. Volume59, Issue 2, March 2016. pp. 195-216.doi.org/10.1111/pala.12221

[MamEv-3] Savage, RJG, & Long, MR (1986).Mammal Evolution: an illustrated guide.New York: Facts on File. p.53.ISBN 0-8160-1194-X.{{cite book}}:CS1 maint: multiple names: authors list (link)

[EoM-4] Gorman, Martyn (1984). Macdonald, D. (ed.).The Encyclopedia of Mammals.New York: Facts on File. pp.766–769.ISBN 0-87196-871-1.

[Grzimek-5] Grzimek, Bernhard.Grzimek's Animal Life Encyclopedia, Volume 10: Mammals I.New York: Van Nostrand Reinhold Company, 1975. Print.

[Animal_clitoris_Jimenez-2013-6] Jiménez, R.; Barrionuevo, F.J.; Burgos, M. (2013). "Natural Exceptions to Normal Gonad Development in Mammals".Sexual Development.7(1–3): 147–162.doi:10.1159/000338768.ISSN 1661-5433.PMID 22626995.S2CID 8721211.

[Animal_clitoris_Carmona-2008-7] Carmona, F. David; Motokawa, Masaharu; Tokita, Masayoshi; Tsuchiya, Kimiyuki; Jiménez, Rafael; Sánchez-Villagra, Marcelo R. (15 May 2008). "The evolution of female mole ovotestes evidences high plasticity of mammalian gonad development".Journal of Experimental Zoology Part B: Molecular and Developmental Evolution.310B(3): 259–266.doi:10.1002/jez.b.21209.ISSN 1552-5007.PMID 18085526.

[Yates&Salazar-Bravo-8] Yates, Terry L.; Jorge Salazar-Bravo (2004). "A Revision OfScapanus latimanus,with the Revalidation of a Species Of Mexican Mole ". In Sánchez-Cordero V.; Medellín R.A. (eds.).Contribuciones Mastozoológicas En Homenaje A Bernardo Villa(PDF).Instituto De Biología e Ins Tituto De Ecología, Unam, México. pp. 479–496.

[9] Castañeda, Sergio Ticul Alvarez; Cortés-Calva, Patrica (2021-05-09)."Revision of moles in the genus Scapanus".THERYA.12(2): 275.doi:10.12933/therya-21-1174.ISSN 2007-3364.S2CID 236583289.

[10] Redescription of the Malaysian Mole as to be a true speciesEuroscaptor malayana^{[permanent dead link]}

[Kawada20122-11] Kawada, S-I.; et al. (2012). "A new species of mole of the genusEuroscaptor(Soricomorpha, Talpidae) from northern Vietnam ".Journal of Mammalogy.93(3): 839–850.doi:10.1644/11-MAMM-A-296.1.

[12] Nicolas, Violaine; Martínez-Vargas, Jessica; Hugot, Jean-Pierre (2017)."Talpa aquitania sp. nov. (Talpidae, Soricomorpha), a new mole species from SW France and N Spain"(PDF).Mammalia.81(6): 641–642.doi:10.1515/MAMMALIA-2017-0057.S2CID 90926022.

[:0-13] Schwermann, Achim H.; He, Kai; Peters, Benjamin J.; Plogschties, Thorsten; Sansalone, Gabriele (2019). "Systematics and macroevolution of extant and fossil scalopine moles (Mammalia, Talpidae)".Palaeontology.62(4): 661–676.doi:10.1111/pala.12422.ISSN 1475-4983.S2CID 134096608.

[14] Hazardous Substances and New Organisms Act 2003 - Schedule 2 Prohibited new organisms,New Zealand Government,retrieved26 January2012

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